integument

integument, in biology, network of features that forms the covering of an organism. The integument delimits the body of the organism, separating it from the environment and protecting it from foreign matter. At the same time it gives communication with the outside, enabling an organism to live in a particular environment.

Among unicellular organisms, such as bacteria and protozoans, the integument corresponds to the cell membrane and any secreted coating that the organism produces. In most invertebrate animals a layer (or layers) of surface (epithelial) cells—often with additional secreted coatings—constitutes the integument. Among the vertebrates the boundary covering—with a variety of derived elements such as scales, feathers, and hair—has assumed the complexity of an organ system, the integumentary system.

The integument is composed of layers that may be of single cell thickness, as in many invertebrates, or multiple cell thickness, as in some invertebrates and all vertebrates. In every case the cells that give rise to the integuments belong to that class of tissue called epithelium, which in most animals is called epidermis. Underlying the epidermis and supplying it with nourishment is the dermis. In addition to the cellular layers, the integument often includes a noncellular coating, or cuticle, that is secreted by the epidermis. Such coatings are found in most invertebrates. The vertebrate skin has generated many kinds of glands and a variety of horny structures, but it lacks coatings.

The wide diversity of integuments among vertebrates further exemplifies the adaptive character of the body covering: from the almost impenetrable shield of an armadillo and the dense furry coat of an Arctic bear to the slimy, scaled covering of a cod and the exceptionally smooth skin of a porpoise. Amphibians and fishes often have mucous glands that lubricate their skins and prevent waterlogging and deterioration. Reptiles have thick, leathery skins that help reduce water loss and serve as an armour against enemies. Birds use their feathers—skin derivatives—to fly and to insulate their bodies. The hairy or furry coats of many terrestrial mammals insulate them, shed water, and provide a dense guard against injury.

Components

The integuments of unicellular organisms comprise the cell membrane and any coating it secretes. Almost all bacteria have an external cell wall that maintains the shape and turgidity of the cell and affords protection. Multicellular invertebrates, however, have a single outer layer of epithelial cells, and these may secrete a variety of surface coatings, ranging from the mucous coat of cnidarians (polyps, sea anemones, jellyfish) to the tough cuticle of insects. The epithelial layer may include cells of several types, such as sensory cells, gland cells, and stinging cells, and the outer surfaces may bear microvilli, cilia, or other fine projections. In addition, the cells may produce excrescences such as bristles, spines, knobs, and ridges.

The firmness of some invertebrate animals, such as annelids (earthworms, marine worms, and leeches) and certain mollusks, depends on the distension by water of the individual cells that form the body wall. In many other forms rigid skeletal materials are deposited either within the cells or on the outer surface. The noncellular coatings of invertebrate integuments are exceedingly varied in composition and extent, and they cut across taxonomic categories. These rigid structures may provide no more than a protective armour, but in the arthropods, including crustaceans, insects, and spiders, a multilayered and hardened integument forms an exoskeleton to which muscles are attached. In the echinoderms, the exoskeleton lies below the epidermis.

Differences among invertebrate groups

The secreted coatings of protozoans exhibit all grades between soft forms (as in Amoeba) and forms with a cuticle that may be proteinaceous (as in Monocystis) or composed of cellulose (as in the plantlike flagellates). Other protozoans have definite shells, composed of protein incorporating various foreign bodies, such as siliceous plates or calcium carbonate (in most foraminiferans), or cellulose (in the resting stages of slime molds). The radiolarians have an internal lattice of silica that is laid down inside the cell—a kind of internal skeleton, or endoskeleton.

Sponges have a simple epithelium, known as the pinacoderm, which both covers the external surfaces and lines the internal waterways. Some sponges deposit needlelike spicules of calcium carbonate in the jelly (mesoglea) beneath this outer epithelium.

In the cnidarians the epidermis provides all the basic features of an integument. It may contain not only epithelial cells, some of which may be contractile, but also gland cells, pigment cells, stinging cells, and sensory cells with projecting hairs. The outer surface often bears flagella or microvilli. The surface secretion may help in capturing food, adhering to substrates, cleaning away settling debris, or providing support and protection. Some hydrozoans produce a horny covering for the polyps, and others have an external skeleton that is calcareous (containing calcium carbonate). Anthozoans show the same diversity. In the common reef-building corals, the calcareous skeleton is secreted by a part of the epidermis that forms a basal disk. This secretory process is continuous, and the polyp raises itself progressively upon a constantly growing stem of calcium carbonate.

The parasitic flukes have a relatively thick integument, which bears many spines and sensory papillae, an apical membrane that is thrown into ridges and pits, and microvilli. The outer part of the integument contains secretory bodies, which are continuously released at the surface to renew the apical membrane. This appears to be a protective device for the parasite related to the immune reaction of the host. Roundworms have a thick, flexible cuticle, with three distinguishable zones, covered by an epicuticle.

Annelids have a thin, horny cuticle pierced by pores through which epidermal glands secrete mucus. In some marine annelids, glands are also present that secrete materials constituting a parchmentlike or calcareous tube within which the worm dwells. Earthworms and leeches secrete cocoons from a specialized epidermis in a region of the body known as the clitellum. A major feature of all annelids except leeches is the possession of bristles, or chaetae, of which there are many varieties. The bulk of each chaeta is secreted by a single cell, though the surrounding lateral cells may contribute materials that bring about its hardening.

The exoskeleton attains its most elaborate forms in the arthropods (for example, crustaceans and insects). The insect epidermis lies on a basement membrane and secretes a tough cuticle, the bulk of which is composed of fibres of a material known as chitin embedded in a matrix of protein. Peripheral to this is an epicuticle. Chitin is a high-molecular-weight polysaccharide containing amino groups. It is synthesized within the epidermis from sugars and amino sugars.

In the integument of caterpillars chitin forms a cuticle that is tough but flexible. But in most arthropods the segments of the body or of the limbs are in the form of rigid plates that form a true exoskeleton linked to adjacent segments by flexible membranes. Such cuticles are hard and may be dark in colour. They are said to be tanned, or sclerotized, and in some species they are also mineralized.

Sclerotization involves the molecular stabilization of the protein chains of the cuticles by establishment of cross-links. Sclerotin, the product of sclerotinization, is a kind of natural plastic. In its horny consistency it closely resembles keratin; both are cross-linked, or polymerized, proteins, but the chemical nature of the linkage is different in the two substances. It is probable that other skeletal proteins in invertebrates, such as the spongin of sponges and the conchiolin of mollusks, are also tanned proteins allied to sclerotin.

In many crustaceans—crabs and lobsters, for example—much of the cuticle is rendered hard by the incorporation of calcareous substances such as aragonite or calcite. But sclerotin is actually harder than calcite, and those parts of crustaceans that need to be of maximum hardness, such as the mandibles and the tips of the claws, are in fact composed of sclerotin.

Besides functioning as a skeleton, the cuticle of terrestrial arthropods must act as a waterproof covering in order to prevent these small animals from drying up. This waterproofing is effected by the secretion of a layer of wax on the surface of the cuticle. Such a wax layer, if exposed in an unprotected state, would be excessively fragile. It is commonly protected by a thin layer of a cementlike substance that is poured over its surface by small dermal glands.

The cuticle of arthropods, pierced by ducts of dermal glands that pour out secretions over the surface, is a living structure; it can produce tactile bristles, pigment-bearing scales, claws, wings, and other structures. In some insects it shows brilliant metallic colours that result from the presence of multiple thin plates or ridges in the cuticle. In order that the arthropod may grow, the old cuticle is shed from time to time after a new and larger cuticle has been laid down beneath it. This process is termed molting, or ecdysis. During the time when the new cuticle is hardening, the arthropod is in a very vulnerable condition.

Molting in insects is hormonally controlled. A molting hormone, known as ecdysone, is mainly a product of the thoracic glands, and its secretion is influenced by a prothoracicotropic (or ecdysiotropic) hormone produced by certain cells of the brain. Larvae also possess a juvenile hormone, which decreases in concentration until the imago (adult) emerges. Crustaceans and spiders possess analogous hormones, though their systems are not identical.

The epidermis of mollusks is capable of a variety of functions. Ciliated epithelium is of particular importance for feeding in bivalves and for the gliding movement of snails. Abundant gland cells secrete mucus, which protects the animal from predators and from desiccation. Complex glands secrete the quinone-tanned proteins of the byssus threads, by which mussels anchor themselves, and of the operculum, with which some sea snails stopper their shells. The secretion of pallial glands enables the date mussel to bore into calcareous rock. Some cephalopods (squids, cuttlefish, octopuses) have luminous glands, although it is disputed whether the luminous material is produced by the epithelium itself or by bacteria. Cephalopods also have pigment cells that can be expanded by muscle contraction and can change colour very rapidly.

The shell of mollusks is secreted by the epithelium of the mantle and consists of an outer layer of the horny substance conchiolin, an intermediate prismatic layer composed of calcite, and a smooth inner layer (the nacreous layer) also composed mainly of calcium carbonate. The first two layers are secreted by a marginal band of cells, so that the shell grows at its outer edge. The nacreous layer is secreted by the general surface of the mantle and is the material of which pearls are formed around foreign bodies introduced into the mantle cavity.

The echinoderms are characterized by a calcareous exoskeleton, which may be a rigid armour, as in echinoids (sea urchins), or of a leathery consistency, as in holothurians (sea cucumbers). The epidermis lies outside of this skeleton. The apical plasma membrane is capable of taking up dissolved organic molecules from the surrounding seawater in amounts that are at least enough for the nourishment of the epidermal layer. Many sea urchins have projecting spines on which the epidermis is worn away to expose the calcareous material.

In all vertebrates the skin has two major layers. The outer, relatively thin epidermis is composed of closely packed cells with little intercellular material; it provides the barrier against attack by chemicals, radiation, or microbes. The underlying dermis (cutis, corium) is thicker and tougher, and its bulk is formed by extracellular materials manufactured by scattered cells. One of its major functions is physical protection. The sensory functions of skin are shared by both epidermis and dermis. The various structures or appendages such as scales, feathers, claws, glands, and hair follicles are derived from either layer or from both. There are considerable differences between the skins of different vertebrate classes, and they are closely related to the environments of the various groups.

The epidermis is the product of the deepest layer of its cells, those that lie immediately over the dermis. From this generative layer, known as the stratum germinativum, cells move outward and become progressively flattened. The surface cells of terrestrial vertebrates, mere remnants of once living cells, are scaly and compressed; they constitute the horny layer, or stratum corneum. The cell fragments of the stratum corneum are composed largely of keratin, a tough insoluble protein. In most land vertebrates the stratum corneum is shed or molted, either periodically and in large fragments or sheets, as in reptiles, or continuously in small patches or scales, as in mammals.

The dermis, which is best developed in mammals, consists largely of fibrous connective tissue (composed of collagen fibres), blood and lymph vessels, smooth muscle cells, and nerve endings. It gives rise to so-called membrane bones—the bony scales of fishes, the bony plates in certain reptiles and mammals, and the membrane bones of the vertebrate skull. Through its blood network the dermis supplies nourishment to the overlying epidermis.

Skin derivatives and appendages

Among the notable changes that have taken place during the course of evolution is the development in vertebrates of a variety of glands, pigmentary structures, scales, claws, nails, horns, feathers, and hairs as adaptations to their changing environments.

Skin glands

The glands of the skin are all exocrine, that is, they secrete their products, usually through ducts, to the epidermal surface. They may be unicellular, as are the goblet cells of fishes, or multicellular, as are the sweat glands of humans. Some multicellular glands are tubular and extrude their secretion into a central space or lumen; some, like the oil-producing sebaceous glands of mammals, form their product by complete breakdown of the cells, a method of secretion known as holocrine. Glands may consist of tubes or sacs, and they may be singular, clustered, or branched; some even contain units of more than one type. They may secrete their product continuously, periodically, or only once.

Mucous glands secrete a protein called mucin, which with water forms the substance known as mucus; this slimy material serves to lubricate the body, thus lessening friction and aiding locomotion in swimming animals. Serous glands produce a watery secretion; sweat glands of mammals are of this type. Sebaceous glands secrete oil, ceruminous glands secrete wax, mammary glands secrete milk, poison glands secrete various toxins, and scent glands secrete a variety of odoriferous substances. Further, certain epidermal glands may be modified into light-producing structures called photophores, seen in the skin of many deep-sea fishes.

Pigment cells

In fishes, pigment is produced in branched cells known as chromatophores, which can be found in both epidermis and dermis. Rapid colour change, by which some fishes can adapt to a change of background, is brought about by redistribution of the pigment within the cell boundaries. Slow, long-term changes involve alterations in the numbers of cells or in the amount of pigment they contain.

Chromatophores are also present in amphibians and reptiles, but not in birds or mammals, which possess pigment cells called melanocytes. Melanocytes are found mainly in the epidermis, though they occur elsewhere. They also are branched, or dendritic, and their dendrites are used to transfer pigment granules to adjacent epidermal cells. A number of different pigments are produced in the different vertebrate groups, but in mammals only brown eumelanin and yellow or red phaeomelanin are important. Pigment cells—chromatophores and melanocytes alike—are influenced by melanocyte-stimulating hormones of the pituitary.

Epidermal scales

Epidermal scales are horny, tough extensions of the stratum corneum. Well developed in reptiles, they are also common on exposed skin in birds and mammals. Such scales are periodically molted or shed gradually along with the rest of the stratum corneum. Epidermal scales are absent in fishes, but dermal, or bony, scales are abundant. Clawlike epidermal scales are present in certain amphibians, including a few toads, certain burrowing, wormlike caecilians, and the salamander Hynobius. The so-called horns of the horned lizard are specialized epidermal scales; and the rattle of rattlesnakes is a series of dried scales loosely attached to each other, the last one always remaining despite molting of the rest of the stratum corneum. Epidermal scales cover the bony scales of the carapace (top) and plastron (bottom) of turtles’ shells. The beak of turtles is composed of a modified epidermal scale covering the jawbone.

Bills of birds are similarly constructed. In birds, epidermal scales are confined to the lower legs, feet, and base of the bill. The spurs of some birds are bony projections covered with a scalelike sheath. The skin of the webs in aquatic birds is also scaly. In mammals, except for a few cases, epidermal scales are largely restricted to the tails and paws. The overlapping horny plates of the pangolin are modified epidermal scales.

In many animals, hardened corneal growths occur at the end of the digits, growing parallel to the skin surface. True claws—found in reptiles, birds, and mammals—consist of a dorsal scalelike plate (unguis) covering a ventral plate (subunguis), the whole capping the bony tip of a digit. Nails—found only in mammals—consist of a broad and flattened unguis, with the subunguis reduced to a vestige under the outer tip. Hooves, the characteristic feature of the hoofed mammals, or ungulates, are exaggerated nails, with the unguis curved all around the end of the digit and surrounding the subunguis.

Horns are hardened corneal projections of several types. Except for certain lizards, horns are found only in mammals. The keratin fibre horn is unique to the rhinoceros. It consists of a cone of keratinized cells that grows from an epidermis covering a cluster of dermal bumps (papillae). The fibres, somewhat resembling thick hair, grow from the papillae, and cells between the papillae produce a cement that binds the fibres together.

Hollow horns are found in cattle, sheep, buffalo, goats, and other ruminants. In certain species only the males display them. Such horns consist of an extension of the frontal bone, a permanent part of the cranium covered by a horny layer. The horn of the pronghorn antelope is unique in that the horny covering is shed periodically and a new one is formed from the epidermis that persists over the bony extension.

Antlers, which are characteristic features of the deer family, are not integumentary derivatives at all. Fully developed antlers are solid bone, without any epidermal covering. The young antlers, however, are covered with skin having a velvety appearance. When the antler is fully developed, the drier skin cracks and is rubbed off by the animal. Antlers in giraffes are small and remain permanently covered.

Feathers and hair

Birds and mammals display remarkable elaborations of the epidermis in the form of feathers and hair, respectively. These distinctive features are dealt with below (see bird; mammal).

Dermal derivatives

Dermal scales are found almost exclusively in fishes and some reptiles. They are bony plates that fit closely together or overlap and form the dermal skeleton. Highly developed dermal scales are seen in turtles, where the bony plates form a rigid dermal skeleton that is attached to the true skeleton. In other reptiles, dermal scales are small and localized on parts of the body, as in crocodilians, certain lizards, and a few snakes.

Birds lack dermal scales, and only a single living mammal—the armadillo—displays them. Associated with the evolutionary tendency toward elaboration of epidermal extensions in birds and mammals, there has been a corresponding reduction in dermal derivatives. The membrane bones of the skull, the mandible (lower jaw), and the clavicles (collarbones) are the remaining vestiges of dermal plates in these groups.

Variations among vertebrates

The vertebrates belong to the phylum Chordata and are closely related to a small, fishlike, almost transparent invertebrate called amphioxus. Amphioxus represents chordate integument at its simplest: an epidermis, consisting of one layer of columnar or cuboidal epithelial cells and scattered mucous cells, covered by a thin cuticle, and a thin dermis of soft connective tissue. Beginning with the simplest vertebrates, the cyclostomes (lampreys and hagfishes), the integument becomes complex and pigmented; in successive evolutionary stages a wide array of derivatives appears among the various classes of vertebrates.

In the lamprey the surface of the skin is smooth, with no scales. The epidermis consists of several cell layers that actively secrete a thin cuticle. Gland cells that produce slime are mixed with the epidermal cells, as in most aquatic vertebrates. The dermis is a thin layer of connective tissue fibres interwoven with blood vessels, nerves, muscle fibres, and chromatophores.

Scales and scale configurations of representative bony and cartilaginous fishes.Encyclopædia Britannica, Inc.Fishes have a more or less smooth, flexible skin dotted with various kinds of glands, both unicellular and multicellular. Mucus-secreting glands are especially abundant. Poison glands, which occur in the skin of many cartilaginous fishes and some bony fishes, are frequently associated with spines on the fins, tail, and gill covers. Photophores, light-emitting organs found especially in deep-sea forms, may be modified mucous glands. They may be used as camouflage or to permit recognition, either for repulsion to delimit territory or for attraction in courtship.

Also formed within the skin of many fishes are the skeletal elements known as scales (Figure 1: Fish scale development.Encyclopædia Britannica, Inc.). They may be divided into several types on the basis of composition and structure. Cosmoid scales, characteristic of extinct lungfishes and not found in any fishes today, are similar to the ganoid scales of living species. Placoid scales (or denticles) are spiny, toothlike projections seen only in cartilaginous fishes. Ganoid scales, sometimes considered a modification of the placoid type, are chiefly bony but are covered with an enamel-like substance called ganoin. These rather thick scales, present in some primitive bony fishes, are well developed in the gars.

Cycloid scales appear to be the inner layer of ganoid or cosmoid scales. Found in carps and similar fishes, they are thin, large, round or oval, and arranged in an overlapping pattern; growth rings are evident on the free edges. Ctenoid scales are similar to cycloid, except that they have spines or comblike teeth along their free edges; these scales are characteristic of the higher bony fishes—perches and sunfishes, for example. Some fishes, such as catfishes and some eels, have no scales.

Among the cartilaginous fishes, sharks have a very tough skin. Scattered over it are denticles, each with a pulp cavity, around the edge of which is a layer of odontoblasts. These cells secrete the dentine, or calcareous material, of the scale. Outside the dentine is the enamel, secreted by the overlying ectoderm. When the denticles pierce through the ectoderm, no more enamel can be added.

The dominant modern fishes, teleosts, are characterized by bony scales covered with skin. The epithelium of a trout’s epidermis provides the animal with an inert covering of keratin. The scales lie in the dermis as thin, overlapping plates with the exposed part bearing the pigment cells. The scale is deposited in a series of annual rings, since its growth occurs rapidly in spring and summer and rarely in winter.

Most modern amphibians lack horny scales or other protective devices. An exception is seen in the caecilians, a small group that has fishlike scales similar to those possessed by ancient and extinct forms. The amphibian epidermis has five to seven layers of cells formed from a basal stratum germinativum. At the skin surface, in contact with the external environment, the cells are keratinized to form a stratum corneum, which is best developed in amphibians that spend most of their time on land. The cells of this horny layer are not continuously shed but are periodically molted in sheets. Molting is controlled by the pituitary and thyroid glands but is unaffected by sex hormones. The wartiness of toads results from local thickenings.

Some amphibian families have disklike pads on their digits for adherence to underlying surfaces. During the breeding season the males of anurans (frogs and toads) and urodeles (salamanders and newts) develop nuptial pads on some digits of the forelimbs, which facilitate firm gripping of the females; the pads are induced to form by androgenic (male) hormones.

The dermis is two-layered, having an outer and looser stratum spongiosum and an inner stratum compactum. Although some amphibians have external gills or internal lungs, for many the skin is a vital respiratory organ, and the dermis is richly supplied with blood vessels and lymph spaces. Chromatophores are located just below the junction of the dermis with the epidermis. The numerous mucous and poison glands originate from nests of epidermal cells that grow down into the dermis.

Reptiles

In the evolutionary sense, reptiles are the first truly terrestrial vertebrates, since they have dispensed with an aqueous environment for their larval development. Their main problem is to prevent desiccation by water loss through the skin. This is solved by the possession of a thick stratum corneum in which waxes are arranged in membranelike layers between the keratinized cells. Reptilian scales are overlapping folds of skin, each scale having an outer surface, an inner surface, and a hinge region. All the epidermal and dermal surfaces of each scale are continuous with those of the next scale.

The cornified part of the epidermis is strengthened by a stiff material, beta keratin, which is present in place of or in addition to pliable alpha keratin. In crocodiles and many turtles the outer scale surface consists of beta keratin only, while the hinge region contains only alpha keratin. In lizards and snakes, however, both keratins form continuous layers, the alpha keratin lying below the beta keratin. In crocodiles and turtles there is continuous cell division in the stratum germinativum and exfoliation of cells at the skin surface. In snakes and lizards the germinal layer forms a complete new epidermal surface before the whole of the old cornified epidermis is sloughed, either in a single sheet or in portions.

The shape and size of the scales vary in the different families and with the mode of life. Maximum flexibility of the skin is achieved in some forms by reduction of the scales to small, nonoverlapping granules. Among desert dwellers there is a tendency for some scales, particularly those on the head and tail, to be enlarged to form spines. Burrowing and secretive forms have a slippery body surface because of the presence of smooth, highly polished scales. The skin is often reinforced by bony plates, which lie beneath the superficial scales (though corresponding with them in size and shape); these plates may form a continuous protective armour. Other defensive, or sometimes offensive, devices associated with the skin and scales are the occasional development of horns or fringing folds that break up the animal’s outline and colouring.

The colours of reptiles are produced by both melanocytes in the epidermis and three types of chromatophores in the dermis: melanophores, which contain melanin; xanthophores, which contain yellow pigments; and iridophores, which contain reflecting platelets of colourless guanine. The pattern may be fixed, for concealment by camouflage, or the chromatophores may provide for rapid colour change.

Reptilian skin possesses glands, but they are usually small. Most are holocrine; some are tubular. Lizards and snakes have small glands that are related to the sloughing cycle, and all groups of reptiles appear to communicate by scent glands. For example, chelonians (turtles and tortoises) have glands in the throat, inguinal, and axillary regions, and snakes have saclike scent glands at the base of the tail.

Birds

The avian epidermis is thin, delicate, and clothed in feathers, except on the obviously naked areas of the legs, feet, beak, comb, and wattle. On the legs and feet, and sometimes elsewhere, the cornified layer is thickened to form scales of several types. The dermis, also thin, consists mostly of a network of connective tissue fibres and muscle fibres that help to adjust the feathers. In larger birds, such as the ostrich, the skin is thick enough to allow it to be processed into leather. The scales resemble those of reptiles in possessing layers containing beta keratin and alpha keratin.

Feathers, which consist of beta keratin, are considered to have evolved from reptilian scales (Figure 2: Development of a typical down feather. The epidermal ridges give rise to the barbs of the feather.Encyclopædia Britannica, Inc.). They are periodically molted, and other keratinized structures such as the bill and claws may be molted as well. Pigment is primarily restricted to feathers and scales. Specialized nerve endings are present throughout the skin. Various holocrine and tubular glands have been observed, but nearly all are small and inconspicuous. The exception is the holocrine uropygial gland, or preen gland, which is located on the back just in front of the tail and secretes oil for grooming the feathers. It is largest in aquatic birds.

Feathers are unique to birds. Those of adults are admirably engineered to be lightweight yet strong. They are of three basic types, each associated with certain functions. Contour feathers (including the flight and tail feathers) define the body outline and serve as aerodynamic devices; filoplumes (hair feathers) and plumules (down feathers) are used principally as insulation, to conserve body heat. Colours and patterns in feathers serve as protective coloration or for sexual display.

In most birds contour feathers are not uniformly distributed over the surface of the body but are arranged in feather tracts (pterylae) separated from one another by regions of almost naked skin (apteria). The only exceptions are the ostrichlike birds, the penguins, and the South American screamers, in which the even distribution of plumage has probably been secondarily acquired. Feather tracts differ in arrangement in different species and hence are useful in the classification of birds.

The wing tract includes the flight feathers proper (remiges) and their coverts (tectrices). The remiges include the primaries, arising from the “hand” and digits and attached to the hand’s skeleton; the secondaries, arising from the forewing and attached to the ulna; and the tertials (when present), arising from the upper wing and attached to the humerus. The tectrices cover the bases of the remiges, overlapping and decreasing in size toward the leading edge of the wing.

The spinal (dorsal) tract extends the whole length of the bird, excepting the head, along and on both sides of the spinal column. In gallinaceous birds this tract may be subdivided from front to back (though not separated by apteria) into the regions of the hackle, the cape, the back, and the saddle. Each region is distinguished by the form and pattern of its constituent feathers.

On the ventral surface of the bird are paired breast tracts, with a ventral tract between them. The tail tract includes the tail feathers (rectrices) and their coverts. Other tracts cover the head, base of the wings, and legs.

A contour feather of an adult bird tends to be almost bilaterally symmetrical. It consists of a tapering central shaft, the rachis, to which are attached a large number of tapering parallel barbs. These in turn carry many minute elongated barbules on both their distal and proximal faces. The distal barbules bear tiny hooklets (hamuli) that fit into grooves on the proximal barbules of the next higher barb. In this way the barbules overlap and interlock to form the coherent web, or vane, of the feather. Barbules in the basal portions of feathers are long, delicate threads and do not bind successive barbs together; consequently, this part of the feather is fluffy.

The filoplumes, which arise at the bases of contour feathers, are inconspicuous hairlike feathers bearing a small tuft of barbs at their apexes. Filoplumes appear to be present in all birds, but only in certain species do they project beyond the contour feathers—on the thighs of cormorants, for example.

Plumules are present in young birds before they develop the adult plumage. In adults the plumules are generally scant and are concealed by contour feathers; however, in many birds, such as gulls and ducks, they form a thick, insulating undercovering comparable to the underfur of seals. Their barbs do not form coherent vanes but are long, loose, soft, and fluffy. Their structure is much simplified, and a rachis may be entirely lacking. In herons and some hawks the tips of the plumules disintegrate into a fine scaly powder that becomes distributed over the plumage, providing protection against wetting and giving it a peculiar sheen; accordingly, these specialized down feathers are called powder down.

Feathers get their colours from a number of pigments. Melanin is responsible for black, gray, brown, and related tints; yellow or reddish brown granules of phaeomelanin and dark brown granules of eumelanin are transferred to the epithelial cells of the feather from melanocytes. Some feathers are coloured bright yellow, vivid red, green, violet, or blue by carotenoids and other rare pigments. Cosmetic coloration of the feathers by the secretion of the preen gland is exploited by pelicans. Not all coloration requires pigments. The striking white of sea gulls and swans is a “structural colour” produced by the reflection of light by irregularly distributed air-filled cavities. Blue, green, and violet can also be structurally produced, as, for example, in kingfishers and parrots.

Mammals

An important distinguishing character of mammals is their hair. They also possess many other horny derivatives of the epidermis, including nails, claws, hooves, quills, and horns. All mammalian hard keratin, as well as the soft keratin of the stratum corneum, is of the alpha type. Bony dermal plates are found in the armadillo. Antlers, too, are made of bone and derived from the dermis, but they have an epidermal covering—the velvet—when newly grown.

Skin structure

The mammalian epidermis has several layers of cells, known as keratinocytes, which arise by cell division in a basal stratum germinativum. This rests on a basement membrane closely anchored to the surface of the dermis. Newly formed cells move outward, and at first form part of the prickle cell layer (stratum spinosum), in which they are knit together by plaquelike structures called desmosomes. Next they move through a granular layer (stratum granulosum), in which they become laden with keratohyalin, a granular component of keratin. Finally the cells flatten, lose their nuclei, and form the stratum corneum. The dead cells at the skin surface are ultimately sloughed, or desquamated. In thick, glabrous skin lacking hair follicles, such as that on human palms and soles, a clear layer, called the stratum lucidum, can be distinguished between the stratum granulosum and the stratum corneum.

The important barrier to outward loss of water or inward passage of chemicals lies in a compact zone of the lower stratum corneum. There the spaces between the layers of the cornified cells are tightly packed with lipid (waxy) platelets that have been produced inside so-called membrane coating granules within the underlying epidermal cells. As well as the clear horizontal stratification of the epidermis, a vertical organization is also apparent, at least in nonglabrous skin, in the sense that the ascending keratinizing cells appear to form regular columns.

In the basal layer, groups of keratinocytes are each associated with a single dendritic (branching) pigment cell to form “epidermal melanocyte units.” In addition to keratinocytes and melanocytes, the mammalian epidermis contains two other cell types: Merkel cells and Langerhans cells. Merkel cells form parts of sensory structures. Langerhans cells are dendritic but unpigmented and are found nearer the skin surface than melanocytes. After a century of question about their purpose, it is now clear that they have a vital immunologic function.

The dermis forms the bulk of the mammalian skin. It is composed of an association of connective tissue fibres, mainly collagen, with a ground substance of mucopolysaccharide materials (glycosaminoglycans), which can hold a quantity of water in its domain. Two regions can be distinguished—an outer papillary layer and an inner reticular layer. The papillary layer is so called by reason of the numerous microscopic papillae that rise into the epidermis, especially in areas of wear or friction on the skin. These papillae, not to be confused with the “dermal papillae” of the hair follicles (see below), are arranged in definite patterns beneath epidermal ridges. In humans these external ridges are responsible for the fingerprints, or dermatoglyphs. The reticular layer has denser collagen than the papillary layer, and it houses the various skin glands, vessels, muscle cells, and nerve endings.

In evolution, the overriding importance of hair is to insulate the warm-blooded mammals against heat loss. Hairs have other uses, however. Their function as sensory organs may, indeed, predate their role in protection from cold. Large stiff hairs (vibrissae), variously called whiskers, sensory hairs, tactile hairs, feelers, and sinus hairs, are found in all mammals except humans and are immensely helpful to night-prowling animals. Vibrissae are part of a highly specialized structure that contains a mass of erectile tissue and a rich sensory nerve supply. These specialized hairs are few in number, their distribution being confined chiefly to the lips, cheeks, and nostrils and around the eyes; they occur elsewhere only occasionally. Human eyelashes consist of sensory hairs that cause reflex shutting of the eyelid when a speck of dust hits them.

Hair may also be concerned in sexual or social communication, either by forming visible structures, like the mane of the lion or the human beard, or by disseminating the product of scent glands, as in the ventral gland of gerbils or the human axillary organ. Hair is important as well in determining the coloration and pattern of the mammalian coat, serving either as camouflage or as a means of calling attention to the animal or a specific part of its body.

In essence, each hair is a cylinder of compacted and keratinized cells growing from a pit in the skin—the hair follicle. The follicle consists mainly of a tubular indentation of the epidermis that fits over a small stud of dermis—the dermal papilla—at its base. Indeed, it is formed in the embryo by just such as interaction between its constituents, the epidermis growing inward as a peg that ultimately invests a small group of dermal cells.

The epidermal components of an active hair follicle consist of an outer layer of polyhedral cells, forming the outer root sheath, and an inner horny stratum, the inner root sheath. This inner sheath is composed of three layers, known respectively as Henle’s layer (the outermost), consisting of horny, fibrous, oblong cells; Huxley’s layer, with polyhedral, nucleated cells containing pigment granules; and the cuticle of the root sheath, having a layer of downwardly imbricate scales (overlapping like roof tiles) that fit over the upwardly imbricate scales of the hair proper. The outer root sheath is surrounded by connective tissue. This consists internally of a vascular layer separated from the root sheath by a basement membrane—the hyaline layer of the follicle. Externally, the tissue has a more open texture corresponding to the deeper part of the dermis that contains the larger branches of the arteries and veins.

A small muscle, the arrector pili, is attached to each hair follicle, with the exception of the small follicles that produce only fine vellus hairs. If this muscle contracts, the hair becomes more erect and the follicle is dragged upward. This creates a protuberance on the skin surface, producing the temporarily roughened condition that is popularly called gooseflesh.

The hair shaft is composed chiefly of a pigmented, horny, fibrous material, which consists of long, tapering fibrillar cells that have become closely impacted. Externally, this so-called cortex is covered by a delicate layer of imbricated scales forming the cuticle. In many hairs the centre of the shaft is occupied by a medulla, which frequently contains minute air bubbles, giving it a dark appearance. The medullary cells tend to be grouped along the central axis of the hair as a core, continuous or interrupted, of single, double, or multiple columns.

The cuticular scales of mammalian hairs are predominantly of the overlapping, imbricate type, with edges that are rounded, minutely notched, or flattened. They vary in size, shape, and edge structure and are distinctive for each species. Among the higher primates, for example, those of chimpanzees are slightly oval, those of gorillas and humans have shallowly notched edges, and those of orangutans have edges that are deeply notched.

In many deer the cortical substance can hardly be distinguished; almost the entire hair appears to be composed of thin-walled polygonal cells. In the peccary the cortical envelope sends radial projections inward, the spaces between being occupied by medullary substance; and this, on a large scale, is the structure of the porcupine’squills.

One of the most remarkable mammalian hairs is that of the Australian duckbill, or platypus, where the lower portion of the shaft is slender and woollike, while the free end terminates as a flattened, spear-shaped, pigmented hair with broad imbricate scales. In the three-toed sloth a microscopic alga grows between the cuticular scales of the hairs and appears to be symbiotic; its presence gives a curious greenish gray hue to the coat of the sloth and helps to disguise the animal among the trees.

The activity of hair follicles is cyclic. After an active period (known as anagen), the follicle passes through a short transition phase (catagen) to enter a resting phase (telogen). In this process, cell division ceases, and the dermal papilla is released from the epidermal matrix, which becomes reduced to a small, inactive, secondary germ. The base of the hair expands and becomes keratinized to form a “club,” which is held in the follicle until the next cycle begins. A new period of anagen starts with cell proliferation of the secondary germ, which then extends inward to reinvest the dermal papilla. After the new hair is formed, the old club hair is shed, or molted. The events of early anagen are, in effect, a reenactment of the early development of the hair follicle.

The final length of any hair depends mainly on the duration of anagen and varies between body sites and from animal to animal. Hairs on the back of a rat take three weeks to grow fully, whereas the follicles on the human scalp may be continuously active for three years or more.

The cyclic activity of hair follicles is the mechanism by which mammals molt; it thus enables animals to alter their coats as they grow or as they adjust to changing temperature-control or camouflage requirements. In some mammals molting takes place in a pattern, so that the follicles act in synchrony in a particular area of the body. In the human scalp the follicles are out of step with each other, and there is continuous loss of club hairs.

Glands

The skin glands of mammals are of three major types. Associated with hair follicles are oil-secreting sebaceous glands as well as tubular glands, which produce an aqueous secretion. Sebaceous glands are termed holocrine because their secretion involves complete disintegration of their cells, which are constantly replaced. Tubular, or merocrine, glands extrude their secretion into a central lumen. The tubular glands of the hair follicle are usually classified as apocrine because it is believed that, in some glands at least, secretion involves a breaking off of part of the gland cells. A second type of merocrine gland, not associated with hair follicles, is termed eccrine because the cells remain intact during secretion. Eccrine glands occur in hairy skin only in humans and some primates; but the footpad glands, which increase friction and thus prevent slipping in many mammalian species, are of a similar type.

A major function of skin glands is the production of odours for sexual or social communication. Many species in all but a few mammalian orders have specialized aggregations of glandular units for this purpose. These occur in almost every area of the body. Some, like the chin and anal glands of the rabbit, contain only tubular units; others, like the abdominal gland of the gerbil, are purely sebaceous; still others, like the side glands of shrews, contain batteries of both holocrine and tubular units.

In some large mammals an important function of merocrine glands is temperature control. Horses and cattle, for example, have apocrine glands for this purpose, but the superbly effective cooling system of humans is served by eccrine sweat glands.

Embryology and evolution

The skin of vertebrates begins to form early in embryonic development, from a superficial germ layer, the ectoderm. The middle germ layer, or mesoderm, proliferates cells rapidly from segmental building blocks, called somites; these cells then migrate in order to lie directly under the outer ectodermal covering. These two embryonic layers—ectoderm and mesoderm—ultimately give rise to the adult skin; the ectoderm produces the epidermis and its derivatives, and the mesoderm produces the dermis.

The human fetus, at least, produces a specialized temporary embryonic skin, known as the periderm. For much of the second trimester of gestation, the periderm consists of cells with projecting globules covered with small protrusions, or microvilli. These cells are subsequently sloughed off as the stratum corneum is formed underneath them.

Differentiation of embryonic tissues proceeds rapidly during the early course of development, and much of what will become adult skin structures—including the glands and appendages—is laid down before the animal is born, often in a latent stage, to resume development later.

As a surface constantly exposed to the environment, the epidermis has undergone more adaptive changes during evolution than any other portion of the skin. Ancestral vertebrates, aquatic and fishlike, were buffeted by water, which kept the living surfaces moist.

The movement to land was gradual and fraught with risk. Amphibians were among the first vertebrates to explore the terrestrial environment. Many evolved a semiaquatic lifestyle, exploiting the land for most of their activities but returning to the water for reproduction. Some remained entirely aquatic, and others adapted to a strictly terrestrial life. Their epidermises reflected such habits: aquatic amphibians developed a thin, slimy, dull skin densely covered with mucous glands; terrestrial forms acquired a thicker, horny, heavily pigmented skin dotted with poison glands.

The reptiles became even more independent of the water. Their skins grew tough, horny, and dry and sometimes received bony contributions from the dermis. Birds evolved a loose, dry skin covered with feathers for insulation and for airfoils and water foils. Finally, mammals adopted a dry, elastic skin, more or less covered with hair. The range of mammalian skin, from smooth (glabrous), as in the cetaceans (whales, dolphins, and porpoises), to densely hairy, as in Arctic bears, is associated with the dispersion of mammals into a wide range of habitats.

Biodynamics

The vertebrate skin—despite its variety—serves the two common functions of protection from, and communication with, the environment. In all land vertebrates the uppermost layers of the skin are dead, but the dermis is richly endowed with living tissue that can respond rapidly to change. A variety of nerve endings constantly report current conditions, and the body makes continuous adjustments in response.

It has been said that the skin is the largest and most versatile organ of the animal body. It shields against injury, against foreign matter and disease organisms, and against potentially harmful rays of the Sun. It also regulates internal body temperature through its insulating ability and its influence on the blood flow. Further, it embodies the sense of touch and adorns the body. Its contours, colour, patterns, and composition aid in species recognition and sexual attraction.

The effectiveness of the skin as a barrier, however, is not complete. Noxious substances that can gain entry evoke an immune response, and the dermis reddens with the rush of blood to the site. Heat also causes expansion of the dermal blood vessels—and in humans and in horses stimulates the sweat glands to heightened activity—thus increasing the loss of body heat. Conversely, cold causes contraction of the vessels and initiates shivering, thereby conserving heat in the first instance and generating it in the second.

The skin is host to a number of microorganisms, especially bacteria and fungi. It is, however, an unstable environment for this population, which lives on the dead epidermal surface that is periodically sloughed off. A normal microcosm exists on most epidermal surfaces. Over the course of evolution an alliance has been established between the skin biota and the epidermal “host,” which tends to stabilize the surface; anything that disrupts the skin biota encourages an imbalance and a potential flare-up of certain microorganisms over others.